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New genera and species of deep-sea Goniasteridae (Asteroidea) from the North Pacific
Abstract
The North Pacific is a vast region, and undiscovered taxa from deep-sea habitats (>200 m), especially from below 1000 m continue to be described based on in situ and museum collections. The Goniasteridae is the most diverse of Asteroidea and further discoveries continue to be made. Here, one new genus and seven new species are described and novel occurrences recorded. In situ observations from R/V Nautilus and MBARI permit further insight into goniasterid ecology and morphology in deep-sea settings.
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Background
Our knowledge of the benthic fauna at lower bathyal to abyssal (LBA, > 2000 m) depths off Eastern Australia was very limited with only a few samples having been collected from these habitats over the last 150 years. In May–June 2017, the IN2017_V03 expedition of the RV Investigator sampled LBA benthic communities along the lower slope and abyss of Australia’s eastern margin from off mid-Tasmania (42°S) to the Coral Sea (23°S), with particular emphasis on describing and analysing patterns of biodiversity that occur within a newly declared network of offshore marine parks.
Methods
The study design was to deploy a 4 m (metal) beam trawl and Brenke sled to collect samples on soft sediment substrata at the target seafloor depths of 2500 and 4000 m at every 1.5 degrees of latitude along the western boundary of the Tasman Sea from 42° to 23°S, traversing seven Australian Marine Parks.
Results
The biological sampling included 35 beam trawls, 28 Brenke sleds, 8 box cores, 20 surface meso-zooplankton tows, and 7 Deep Towed Camera transects. In total, 25,710 specimens were identified to 1084 taxonomic entities, including 847 species-level, 144 genus-level and 69 family-level and 24 higher-level taxa. Of the species-level taxa, only 457 were assigned species-level taxonomic names, which implies that up to 58% of the collected fauna is undescribed. In addition, the ranges of numerous species have been extended to include the western Tasman Sea.
Conclusions
The lower bathyal and abyssal fauna of soft sediment seafloors off eastern Australia has been systematically surveyed for the first time. The resultant collections will provide the foundation for much future ecological, biogeographical, phylogenetic and taxonomic research.
Background: The Great Australian Bight (GAB) comprises the majority of Australia’s southern coastline, but to date its deep water fauna has remained almost unknown. Recent issuing of oil and gas leases in the region has highlighted this lack of baseline biological data and established a pressing need to characterise benthic abyssal fauna.
Methods: From 2013 to 2017, six large-scale systematic surveys of the GAB were conducted from 200 to 5000 m depth, constituting the deepest systematic biological sampling in Australia. Sampling was conducted on soft sediment and hard substrates, both at predetermined depth intervals along north-south transect lines and at sites of interest identified by multibeam sonar.
Results: A total of 66,721 invertebrate specimens were collected, comprising 1267 species, with 401 species (32%) new to science. In addition to the novelty of the fauna, there was a high degree of rarity, with 31% of species known only from single specimens.
Conclusions: In this paper, we provide an annotated checklist of the benthic invertebrate fauna of the deep GAB, supplemented with colour photos of live specimens and commentary on taxonomy, diversity and distributions. This work represents an important addition to knowledge of Australia’s deep sea fauna, and will provide the foundation for further ecological, biogeographical and systematic research
The sea pen Ptilosarcus gurneyi Gray provides a major source of food for at least seven predator species in Puget Sound: the asteroids Hippasteria spinosa, Dermasterias imbricata, Crossaster papposus, and Mediaster aequalis, and the nudibranchs Armina californica, Tritonia festiva, and Hermissenda crassicornis. Ptilosarcus is long-lived (15 yr +), takes 5 or 6 yr to reach sexual maturity, and has a spatially clumped pattern of recruitment. It never grows large enough to avoid predation by four of its predators, and the refuges and escape mechanisms of Ptilosarcus are not sufficient to explain its abundance in the face of this predation. Determination of rates of critical life history processes of species involved in this association, e.g., recruitment, growth, and predation, permitted estimates of the effects of the activities of each species on the others in the association. Particular attention was paid to age structure and to the natural history during the periods shortly following metamorphosis of each species. In dense recruitment patches, 97% of Ptilosarcus mortality occurs during the 1st year because of predation by Hermissenda, Tritonia, and Crossaster. Adult pens are eaten by Hippasteria, Dermasterias, Mediaster, and Armina, which together remove 2 pens/m^2@?yr, or 3.1% from the 10 adult year classes combined. The latter four predator species selectively prey upon the largest pens available. The time required for capture and consumption increases at a slower rate with prey size than does prey biomass. Recruitment of the long-lived food specialist Hippasteria seems to be impaired by the spatially unpredictable and clumped nature of Ptilosarcus recruitment. Although the adults have an opulent food supply, the young Hippasteria require small Ptilosarcus to feed on and rarely find enough. The generalists, which are not dependent upon one prey during their recruitment and are able to maintain themselves without Ptilosarcus, could in theory increase until they have reduced the sea pen populations to below the level needed by the specialists and thus outcompete them. However, predation on the generalists by a higher order predator, the asteroid Solaster dawsoni, removes each year at least 9.4% of the adult Mediaster and 38% of the adult Crossaster, a rate at least as great as, and usually greater than, the rate of recruitment into the populations of adult prey. The characteristic organization of the Ptilosarcus association and the existence of Hippasteria may depend upon S. dawsoni.
Genetic structure and connectivity of populations of the globally distributed and eurybathic sea star Hippasteria phrygiana (Parelius 1768) were studied in 165 individuals sampled from three oceanic regions: the North Pacific Ocean, the South Pacific Ocean (considered to include the adjacent regions of the Southern Ocean and the southern Indian Ocean) and the North Atlantic Ocean. A nuclear gene region (ATP synthase subunit α intron #5, ATPSα) and a mitochondrial gene region (cytochrome oxidase subunit I, COI) were amplified and sequenced. Significant heterogeneity was detected in an AMOVA analysis among the three sampled oceanic regions for COI, but not for ATPSα. Neither gene showed significant genetic heterogeneity within the North Atlantic, as assessed by ΦST values. Significant heterogeneity was detected for COI (but not ATPSα) in the North Pacific, but the converse was true in the South Pacific. Coalescent simulations suggested that the three regions have been diverging with little or no gene flow for the past 50–75,000 years, a time frame that corresponds to the onset of the last glacial period of the Pleistocene. A possible genetic signature of recent population expansion (or non-neutrality) was detected for each gene in the North Pacific, but not in the other two oceanic regions.
A phylogenetic analysis of 13 taxa and 32 characters resulted in a single most parsimonious tree that supports monophyly of the goniasterid (Echinodermata, Asteroidea) genus Circeaster Koehler, 1909 and supports re-establishment of the genus Lydiaster Koehler, 1909. Th e phylogeny supports monophyly of the ingroup, including 10 species, six of which, C. kristinae n. sp., C. helenae n. sp., C. arandae n. sp., C. loisetteae n. sp., C. sandrae n. sp., and C. pullus n. sp., are new. Phylogenetic results support diversifi cation into the Indian, Pacifi c, and Atlantic ocean basins. Th e phylogeny is constrained by a sister taxon with a Cretaceous fossil occurrence and two geologic events, including the closure of the Indonesian seaway and formation of the Panamanian isthmus. Th ese events formed barriers limiting or preventing larval dispersal between the Indian/Pacifi c and the Pacifi c/Atlantic oceans. Larval dispersal through a deep-sea environment was a signifi cant consideration for estimating timing constraints from paleo- environments. Based on fossil constraints, ancestry for the lineage is suggested as early as the Late Cretaceous with subsequent diversifi cation in the Cenozoic. In situ observations of Circeaster perched on bare deep-sea coral skeletons and morphological similarities with other known corallivorous goniasterids suggest important ecological roles in the deep-sea.
Tropical shallow water sea star faunas, especially those of the Indo-West Pacific, are dominated by the order Valvatida. Among sea stars, valvatidans have the best-developed antipredatory devices. Vermeij (1978) found high to low latitudinal increases in antipredatory structures in various in vertebrate groups (e.g., gastropods). The valvatidan occurrences suggest the presence of controls in sea stars similar to those affecting other groups. The Valvatida includes few genera that prey on active, solitary invertebrates, but such habits are common in other orders, and in cooler waters. Protective structures appear to restrict predatory abilities. The importance of sea stars as predators on solitary organisms declines in tropical latitudes, yet sea stars have evolved only limited basic structural variation since their appearance in the Ordovician. Phylogenetic constraints in adaptability appear strong in sea stars because of their evolutionary failure to maintain predatory life habits in shallow tropical waters.
Ptilocrinus amezianeae n. sp. is a new species of stalked crinoid attributed to the family Hyocrinidae. Forty-five specimens were collected from
seamounts north of the Ross Sea, and one specimen from the Kerguelen Plateau at depths ranging from 450 to 1,680m. The collection
from Admiralty and Scott seamounts constitutes the first example of a hyocrinid population known both from in situ photographs
and from numerous collected specimens ranging from small juvenile to large adult. Variation in theca and stalk articulation
characters throughout ontogeny is congruent with the molecular data and indicates that all the specimens examined belong to
a single species. Tegmen and pinnule architecture, brachial arrangement, and stalk articular facets indicate that Ptilocrinus amezianeae n. sp. has close affinities with P. clarki and P. pinnatus from the northeastern Pacific and displays the most derived characters among these three species. Two cases of true arm division
into two unequal branches suggest that Ptilocrinus and Calamocrinus are closely related. The picture and video transects on Admiralty seamount show a patchy distribution of living specimens
with patches of mean density ca. 2.6 individuals m-2. In situ photographs also document predation by a sea urchin and a sea
star on tegmen and proximal arms. The COI gene sequences analyzed in 25 specimens from Admiralty and Scott seamounts display
low pairwise distances, low nucleotidic diversity, and intermediate haplotype diversity. These results, together with disarticulated
ossicles and attachment disks observed on in situ photographs, indicate that the population investigated here is in decline.
KeywordsEchinodermata–Stalked crinoids–Hyocrinidae–
Ptilocrinus
–Antarctica–Seamount–Ross sea–Kerguelen plateau
Between 2015 and 2018, the NOAA ship Okeanos Explorer embarked on the CAPSTONE expedition, that collected video and specimen data from the Hawaiian Islands region and other American territories in the Central and South Pacific Ocean. Ultimately 20 species, 12 of which are newly described, from 14 genera in 3 families are described herein with one further undescribed species of Pythonaster currently in preparation. Okeanos Explorer collected a total of 17 sea star specimens, of which 9 represent new species and 4 new genera. Additionally, one species imaged by Okeanos Explorer during the campaign was discovered in a museum collection is described herein. Supplementary collections from the tropical Pacific by the R/V Nautilus resulted in a further 11th newly described species which had been originally observed by Okeanos Explorer. A total of 21 specimens were involved as part of this effort. Heligmaster n. gen. is the first new genus of the rarely encountered deep-sea Myxasteridae since Fisher (1906). The Goniasteridae, particularly the corallivorous subfamily Hippasterinae were most heavily represented in the collected material. Video imagery was extensive and provided new occurrence data for rarely seen species as well as several new records that likely represent undescribed species. Many of the new species collected were observed preying on of deep-sea octocorals and sponges.
Seastars are some of the most beautiful and intriguing animals living in our oceans. Their shapes, diversity of colours, sizes and surface patterns are unique and amazing. Found from the deep sea to the shallow waters of intertidal areas, these 'stars of the sea' are often encountered by swimmers, snorkellers and divers, or even washed up on shore after storms and wild seas.
This comprehensive guide introduces the shallow water seastars of Australia, how to identify them, where they live, what they eat, when and how they reproduce, and many other fascinating aspects of their biology
Exploratory cruises by the NOAA Ship Okeanos Explorer have resulted in a substantial contribution in our understanding of deep-sea echinoderm biodiversity, biology, and ecology in the North Atlantic. This includes the description and in situ feeding observations of two, new corallivorous goniasterid species, Evoplosoma nizinskiae n. sp. and Sibogaster bathyheuretor n. sp. Significant in situ observations include a synchronous feeding event including multiple goniasterid asteroids and a cidaroid urchin on a large demosponge, providing new data for understanding echinoderm feeding behavior, including agonistic behavior, in deep-sea settings and new, in situ feeding observations for 28 deep-sea species including the myxasterid Pythonaster atlantidis, the korethrasterid Remaster palmatus and the poorly understood hippasterine goniasterids, Gilbertaster caribaea and Sthenaster emmae.
Modern goniasterids are the most numerous of living asteroids in terms of described genera and species and they have important ecological roles from shallow to deep-water marine habitats. Recent MNHN expeditions and historical collections in the USNM have resulted in the discovery of 18 new species, three new genera and multiple new occurrence records from the western Indian Ocean region including Madagascar, Glorioso and Mayotte islands, Walters Shoal, South Africa, and Somalia. This report provides the first significant contribution to knowledge of deep-sea Asteroidea from the Indian Ocean since the late 20th Century. Several deep-sea species, previously known from the North Pacific are now reported from the western Indian Ocean. Gut contents from Stellaster and Ogmaster indicate deposit feeding. Feeding modes of this and other deep-sea species are discussed. Comments are made on fossil members of included taxa. A checklist of Indian Ocean Goniasteridae is also included.
Research cruises by the Monterey Bay Aquarium Research Institute (MBARI) have resulted in the discovery of three new species of asteroids from the lower bathyal/abyssal regions of the North Pacific. A new genus, Bathyceramaster is described to accommodate "Mediaster" elegans Ludwig 1905 and related species. New records and in situ observations are also presented. An identification key and taxonomic account of the deep-sea Goniasteridae known from this region including one new genus and two new species is included. An overview of the genus Sibogaster, including a new widely occurring species, is also included.
A review of high-latitude Goniasteridae south of 50°S is presented, including a key to species, figures and taxonomic descriptions of all species with a brief taxonomic summary for all genera. Nineteen species in 11 genera are reviewed. Era-tosaster jenae nov. gen and sp., Chitonaster trangae nov. sp., and Pillsburiaster calvus nov. sp. are described. Several taxa, including Chitonaster, have not been reviewed since their initial description and revision of the group has clarified taxonomic boundaries between species. The genus Pentoplia is a junior synonym of Chitonaster. Rarely encountered species, synonyms, and range extensions are reported for multiple species.
Sea-stars in the orders Velatida, Spinulosida, Forcipulatida and Brisingida are described for the New Zealand biogeographic region (latitudes 24° to 57° 30′ S, 157° E to 167° W). The area includes the New Zealand Exclusive Economic Zone (EEZ), Lord Howe, Norfolk, and Macquarie Islands, the Kermadec Islands and associated ridges, and deeper basins and trenches. In all, 16 families, 39 genera and 83 species are represented. The order Velatida includes 5 families, 10 genera, and 23 species; Spinulosida includes 2 families, 4 genera, and 14 species; Forcipulatida 4 families, 17 genera, and 31 species; and Brisingida 5 families, 8 genera and 15 species. Families new to the fauna are Myxasteridae (Velatida), Leilasteridae (Spinulosida), and Pedicellasteridae (Forcipulatida). One new genus is described in the family Asteriidae (Forcipulatida). New species are described in the families Pterasteridae and Myxasteridae (Velatida), and Pedicellasteridae, Zoroasteridae, and Asteriidae (Forcipulatida). Species not formally recognised are present in the Pterasteridae, Zoroasteridae, and Hymenodiscididae. The order Forcipulatida dominates the fauna in number of species and recorded occurrences, while the Spinulosida has least species and occurrences. The Velatida have the largest bathymetric range (0-6720 m), followed by the Forcipulatida (0-4405 m). The Spinulosida are decidedly shallower (0-1357 m), while the Brisingida (150-6160 m) are the only order not recorded from shallow water. At least 34 species are endemic, and only 8 are recorded from depths greater than 3000 m. An addendum includes species in the orders Paxillosida (1), and Valvatida (17) not previously recorded locally. They include the genera Patagiaster (Astropectinidae, Paxillosida), Astroceramus and Cladaster (Goniasteridae), two new genera in the families Goniasteridae and Odontasteridae, and new species in the genera Astroceramus, Cladaster, Ceramaster, Pillsburiaster (Goniasteridae), Marginaster (Poraniidae) and Podosphaeraster (Podosphaerasteridae) in the order Valvatida. Also included are Mediaster australiensis H.L. Clark; Pseudarchaster jordani Fisher; and small specimens of Ceramaster which may be two distinct species and a species of Marginaster, though these last three are not formally named. Additional records are listed for Anseropoda aotearoa and Tremaster mirabilis novaecaledoniae. Two species of Hippasteria are included, though from outside the area; one is H. falklandica Fisher, the other is new.
A new species of corallivorous goniasterid, Evoplosoma watlingi n. sp., from Bear Seamount in the North Atlantic is described in conjunction with an overview of other species within the genus Evoplosoma. Hippasteria tasmanica McKnight, 2006 is re-classified within Evoplosoma. In situ images of Atlantic deep-sea Goniasteridae, including Evoplosoma spp., Hippasteria phrygiana and Circeaster americanus, are presented herein. Observations of several Evoplosoma species and the Atlantic C. americanus are documented for the first time, recorded by the R/V Okeanos Explorer. The feeding duration of C. americanus is also documented for the first time.
Asteroids (sea stars) can be important predators in benthic communities and are often present in ecologically important and vulnerable deep-sea coral and sponge habitats. However, explicit studies on the trophic ecology of deep-sea asteroids are rare. We investigated the diets of seven species of deep-sea asteroid from the bathyal zone of Newfoundland and Labrador, eastern Canada. A multifaceted approach including live animal observations, stomach content analysis, and stable isotope analysis revealed the asteroids to be either top predators of megafauna or secondary consumers (mud ingesters, infaunal predators, and suspension feeders). The stable isotope signatures of Ceramaster granularis, Hippasteria phrygiana, and Mediaster bairdi are characteristic of high-level predators, having δ15N values 4.4‰ (more than one trophic level) above Ctenodiscus crispatus, Leptychaster arcticus, Novodinia americana, and Zoroaster fulgens. We present strong evidence that corals and sponges are common food items for two of the predatory species, C. granularis and H. phrygiana. During laboratory feeding trials, live H. phrygiana fed on several species of soft coral and C. granularis fed on sponges. Stomach content analysis of wild-caught individuals revealed sclerites from sea pens (e.g. Pennatula sp.) in the stomachs of both asteroid species; H. phrygiana also contained sclerites from at least two other species of octocoral and siliceous sponge spicules were present in the stomachs of C. granularis. The stomach contents of the secondary consumers contained a range of invertebrate material. Leptychaster arcticus and Ctenodiscus crispatus feed infaunally on bulk sediment and molluscs, Zoroaster fulgens is a generalist infaunal predator, and the brisingid Novodinia americana is a specialist suspension feeder on benthopelagic crustaceans. This study provides a foundation for understanding the ecological roles of bathyal asteroids, and suggests that some species may have the potential to be important modulators of deep-sea benthic communities.
The herbivorous gastropod Tegula funebralis is not highly ranked in a food preference hierarchy of its major predator, the starfish Pisaster ochraceus, and exhibits a persistent broad overlap with it in the rocky intertidal zone at Mukkaw Bay, Washington. Observations on Tegula over a 5-yr period indicate that it settles high intertidally, lives there for 5-6 yr, and then tends to migrate lower into contact with Pisaster. Tegula lays down an annual growth line permitting it to be aged and a growth curve constructed. Analysis of relative growth and reproduction indicates that beyond a certain size (16 mm) large individuals perform less well in the upper than those in the lower intertidal zone. Pisaster consumes 25-28% of the adult Tegula per year in the area of spatial overlap, based on analysis of the age structure of 6-17 yr old Tegula, and by direct estimates of the percentage of the standing crop consumed annually. The relationship between Pisaster and sex ratio, relative energy limitation and reproductive output (fitness) of Tegula is discussed for three subpopulations. It is suggested that the implied results of the interaction is typical of that between a major predator and one of its less preferred prey. The prominent zonation exhibited by preferred prey, the observed intimacy of association of predator and less preferred prey, and the zoogeographic homogeneity of the Pacific rocky coastline community are discussed in relation to three intermeshing ecological processes.
In the course of this work over 3000 specimens were examined and recorded. These came from the two largest collections in New Zealand, NIWA (incorporating the New Zealand Oceanographic Institute) and the Museum of New Zealand. Collections at the Auckland Museum were also reviewed. The order Paxillosida is represented by four families-Luidiidae (1 genus, 5 species), Astropectinidae (8 genera, 24 species), Radiasteridae (1 genus, 2 species), and Porcellanasteridae (5 genera, 7 species). There are eight new species. The order Notomyotida is represented by one family-Benthopectinidae (3 genera, 12 species, none new). Specimens were collected from very shallow tidal areas to nearly 5000 m. A typical specimen is described in detail for each species, and geographic distribution and variations in depth are also discussed. Photographs and drawings of relevant parts are included. In three very common species with many individuals (Astromesites primigenius, Astropecten polyacanthus, and Psilaster acuminatus), a number of specimens were dissected and stomach contents listed. No parasites were observed. Dissected specimens appeared to be in good condition with generally well-developed gonads. Notes on internal anatomy and other features of interest are included where applicable.
Eleven species of goniasterid sea stars collected by the R/V Gerda during a biological survey of the Straits of Florida were studied. Each is fully described and illustrated. A key to all the goniasterid genera in the tropical western Atlantic is provided.Wide variations occur within many of the species, and as a result, many invalid nominal species have been described. Thus, a good deal of taxonomic revision is made. Ceramaster affinis is made a junior synonym of C. grenadensis. The genus Peltaster is delimited and placenta placed in it. Peltaster planus is made a junior synonym of P. placenta. Plinthaster perrieri, P. nitidus, and P. comptus are made junior synonyms of P. dentatus. Lydiaster is made a junior synonym of Circeaster, and Circeaster occidentalis is made a junior synonym of C. americanus.
The previously unsurveyed Admiralty (67°S, 171°E) and Scott Island (67°22′S, 179°55′E) seamounts to the north of the Ross Sea were explored using cameras and physical sampling gear during the New Zealand IPY-CAML research voyage (TAN0802) in February 2008. A striking assemblage dominated by stalked crinoids and brachiopods was found at 580–600 m depths on isolated knolls at the northwest and southeast extremities of Admiralty Seamount. The seabed at these sites was littered with crinoid ossicles, and crinoid stalk bases were conspicuous on exposed rocks, suggesting that these assemblages have persisted for a considerable period of time. The crinoid sites were limited to the isolated knolls but large areas of the flanks of the main seamount were covered by dense populations of suspension-feeding ophiuroids. These assemblages are more similar in structure to those preserved in fossil strata from the Palaeocene and late Eocene than to any extant assemblages yet described from the Antarctic. In comparisons with faunal assemblages on Scott Island seamount, the abundance of stalked crinoids was strongly inversely correlated with the abundance of echinoid, asteroid, and lithodid crab predators and both asteroids and echinoids were photographed feeding on crinoids. These observations are consistent with the prevailing hypotheses that crinoid- and ophiuroid-dominated assemblages, which were widespread in the Palaeocene, were displaced by the radiation of mobile predators, and that conditions in isolated habitats in the Southern Ocean may have acted as refugia, allowing the persistence of archaic benthic assemblages.
The Valvatacea is one the most ecologically important, taxonomically diverse, and widespread groups of post-Palaeozoic (i.e. modern) Asteroidea. Classification within the group has been historically problematic. We present a comprehensively sampled, three-gene (12S, 16S, early-stage histone H3) molecular phylogenetic analysis of the Valvatacea. We include five of the six families within the Paxillosida, the monotypic Notomyotida, and 13 of the 16 families of the living Valvatida. The Solasteridae is removed from the Velatida (Spinulosacea) and joins the Ganeriidae and the Leilasteridae as members of the clade containing the Asterinidae. The Poraniidae is supported as the sister group to the large cluster of Valvatacea. Asteropseids and poraniids are phylogenetically distant, contrary to morphological evidence. Several goniasterid-like ophidiasterids, such as Fromia and Neoferdina are supported as derived goniasterids rather than as Ophidiasteridae. The Benthopectinidae (Notomyotida) are supported as members of the Paxillosida as are two members of the Pseudarchasterinae that have traditionally been considered members of the Goniasteridae. Our data suggest that Antarctic valvataceans may be derived from sister taxa in adjacent regions.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161, 769–788.
It is suggested that local animal species diversity is related to the number of predators in the system and their efficiency in preventing single species from monopolizing some important, limiting, requisite. In the marine rocky intertidal this requisite usually is space. Where predators capable of preventing monopolies are missing, or are experimentally removed, the systems become less diverse. On a local scale, no relationship between latitude (10⚬ to 49⚬ N.) and diversity was found. On a geographic scale, an increased stability of annual production may lead to an increased capacity for systems to support higher-level carnivores. Hence tropical, or other, ecosystems are more diverse, and are characterized by disproportionately more carnivores.
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